Journal of BIOLOGICAL RESEARCHES ISSN: 08526834 | E-ISSN:2337-389X Volume 21| No

Home , Ingerana, Kalophrynus, Limnonectes, Megophrys, Metaphrynella, Philautus

Journal of BIOLOGICAL RESEARCHES ISSN: 08526834 | E-ISSN:2337-389X Volume 21| No. 1| December | 2015

Review Article

SPECIATION AND ZOOGEOGRAPHY OF AMPHIBIAN IN SUNDALAND

Nia Kurniawan1*, Driyana Rike Ahmadlia2, Day Shine Nahari3, Anggun Sausan Firdaus1

1Biology Department, Faculty of Mathematics and Natural Sciences, University of Brawijaya 2Biology Department, Faculty of Sciences and Technology, Islamic State University Maulana Malik Ibrahim 3Chemistry Department, Faculty of Sciences and Technology, Islamic State University Maulana Malik Ibrahim Veteran Street, Malang 65145, East Java, Indonesia

ABSTRACT

Sundaland is an interesting area to be explored based on its geological history, topography, and climate. Sundaland consists of Penisular Malaysia, Sumatra, Borneo, and Java which experienced some emergence and submergence process in the past. During 1981-2015, most of research in Sundaland found that amphibian family in Sundaland was dominated by Bufonidae, Ranidae, Microhylidae, Megophrydae, Rachophoridae, and Dicroglossidae which experienced lot of speciation in its history. Among of 4 major islands in Sundaland, Borneo has the highest number of species diversity, then Peninsular Malaysia, Sumatra, and Java. During those years, Sumatra and Java got least concern by researcher. There- fore, it is suggested for further study to explore more in Sumatra and Java.

Keywords: Sundaland, amphibian, speciation, zoogeography.

INTRODUCTION Amphibian composition in Sundaland is the result of amphibian exchange at Plio-Pleistocene period (Wilting et The history of Sundaland has begun at the Early- al., 2012; Brandon and Jones, 1996). There were several Cenozoic (approximately 65 mya). It was merged with the factors which influenced amphibian composition in southern end of Eurasia continent (Hall, 1996). During Sundaland, they are geological history of Sundaland and Pliocene and Pleistocene period, Sundaland was a large tropical climate which affected on the other geological single landmass. Since the rise of sea level in glacial factors, such as ancient river system and climate. Beside period, Sundaland dispersed into Java, Borneo, Sumatra, that, supervulcanic eruption also gave some influence Malay-Peninsula, and some adjacent islands (Molen- (Woodruff, 2003; Lohman et al., 2008). This chapter will graaff, 1916). discuss about speciation and zoogeography of amphibians Sundaland has dispersed along the equator line with in Sundaland, despite on its geological history and mountains and rivers on it. Sundaland has tropical climate amphibian diversity (especially amphibians).Amphibian and seasonal wind. During glacial Pleistocene, the climate composition in Sundaland is the result of amphibian ex- has predicted to be cooler and drier, except in Sumatra, change at Plio-Pleistocene period (Wilting et al., 2012; West Borneo, and North Borneo which have wet climate. Brandon and Jones, 1996). Most of Sundaland included to monsoon climate area and the rest northern area with equatorial climate. The difference is on its rain cycle in the monsoon climate SPECIATION OF AMPHIBIAN IN region. Climate in Sundaland is fluctuative between SUNDALAND cooler-drier periods and warm-wet periods. Both of the Speciation in Sundaland was predicted has begun periods has influenced the occurrence of tropical forest when the process of land connecting and separating as the area as habitat of Sundaland amphibian in the past. On the result of sea-level fluctuation since Eocene until other side, warmer climate in Sundaland causes it has Pleistocene period (Wilting et al., 2012; Ruedi, 1996). high biodiversity. Warmer climate provide more food and Both of these processes end with allopatric speciation constant temperature in a year. Thereby, Sundaland has which caused by physical barrier (such as topography, high biodiversity than other subtropical regions (Lohman water, and landmass), it leads to the formation of genetic et al., 2011). reproduction barrier. Futuyama (2005) explained in his book “Evolution” that natural selection and genetic drift  Corresponding Author: have caused the divergence in genetic composition. When Nia Kurniawan a region is separated in a long period of time Biology Department, Faculty of Mathematics and Natural Sciences, Brawijaya University (geographically) it will form new species as the result of Jl. Veteran, Malang 65145, East Java, Indonesia allopatric speciation. phone : +6282142452370 Amphibian research in Sundaland has started since e-mail : [email protected] the 18th until 20th century (today) and has completed the database of amphibian. There are 6 families which found

Open Access  www.berkalahayati.org 1 Speciation and Zoogeography of Amphibian in Sundaland

in Sundaland (Bufonidae, Ranidae, Megophrydae, estimated have some species resulted from speciation in Microhylidae, Dicroglossidae, and Rhacophoridae) with their developmental process. Percentage of each family in the high similarity on their characters and high species Sundaland (total species in Sundaland) since 18 century diversity in every island. During the 18th until 20th century was Rhacophoridae 25%, Ranidae 20%, Bufonidae 18%, some studies showed that Sundaland has 301 species of Microhylidae 14%, Megophrydae 12%, and amphibian (from 6 large families) approximately which Dicroglossidae 11% (Figure 1.a). some of them are endemic species. Each family was

Figure 1. a) Percentage of amphibian species based on distributed family in Sundaland; b) Percentage of new species which distributed in Sundaland.

bution

(%) Percentage Distr Species of

Figure 2. Percentage of species distribution in amphibian based on 6 main family (Bufonidae, Ranidae, Megophrydae, Microhylidae, Rhacophoridae, and Dicroglossidae) in 4 major islands (Malay Peninsular, Borneo, Sumatra, and Java) and adjacent island.

Rhacophoridae distributed almost in all 4 major islands (Teynie et al., Rhacophoridae family has the highest species 2010; Streitcher et al., 2014; Harvey et al., 2002). diversity in Sundaland compared with five other families Philautus and Rhacophorus were genera with the (see Figure 1.a). It consists of ±6 genera (Nyctixalus, highest number of densityin Sundaland compared with Phlautus, Polypedates, Rhacophorus, Theloderma, and other genera (Teynie et al., 2010). Rhacophoridae has Chiromantis) with total ±74 of species. Some species in highcharacter similaries among its species member which Rhacophoridae have been predicted as endemic species in predicted as the result of geological condition of Sumatra and Java. Species which predicted as endemic in Sundaland that led to speciation. Philautus ingeri is one Java were Philautus jacobsoni, Philautus pallidipes, of the example speciation results which found at Sarawak Philautus vitiger, and Rhacophorus javanus (Kurniati, (Borneo) and predicted as P. hosei by Dring (1987). 2006), while Sumatra has Rhacophorus achantharrena, Another example is Philautus mjobergi which identified Rhacophorus barisani, Rhacophorus bifasciatus, by Smith (1925) from Borneo, it first known as P. Rhacophorus bengkuluensis, Rhacophorus catamitus. aurifasciatus (Bossuyt and Dubois, 2001). High Rhacophorus, Polypedates, and Theloderma genera have similarityof their morphological characters sometimes wide distribution area in Sundaland, because they are cause misidentification. Morphological similarities of the species were predicted as the result of speciation process,

Journal of BIOLOGICAL RESEARCHES | Volume 21 | Number 1 | December | 2015 2 Kurniawan et al. as described by Darwin that evolution was originated distribution in Malay Peninsula, and Borneo predicted from the ancestor which have changed and developed caused by the geological history of Sundaland. (Coyne, 2009). Ansonia breeds in stocky stream which facilitated by land mass of Riau-Bangka-Karimata arcs and Anambas- Ranidae Natuna arcs. Genetical exchange of Ansonia was Ranidae consists of ±7 genera which spread widely predicted occurred when Borneo and Peninsular Malaysia in Sundaland (Amolops, Huia, Hylarana, Staurois, were separated from Sumatra and Javathen caused Meristogenys, Odorrana, and Rana) with ±61 species. allopatric speciation (Inger and Voris, 2001). Some Rana has the highest species diversity compared with species of Ansonia have been clustered and only other genus and spread in Peninsular Malaysia, Borneo, distributed in Peninsular Malaysia or Borneo. The Sumatra, and Java. As the genus with the highest species predicted scenario for this condition is the vicariance diversity and the widest distribution, Rana has some event, the increasing of sea level that caused the species which resulted from speciation (Lim and Leong, separation of area distribution between Peninsular 2004; Tjong et al., 2012). For example Rana siberu which Malaysia and Borneo which occurred at Early-Miocene formerly distributed in Sumatra, including Aceh, (24 – 13 Mya) (Bohlen et al., 2011; Woodruff, 2003). Bengkulu, and West Sumatra (Iskandar and Coljin, 2002), Ansonia then fragmented by mountain and sea barriers species also found in Peninsular Malaysia recently (Lim which lead to the natural selection and genetic drift. and Leong, 2004). Rana siberu firstly described as R. signata complex Microhylidae because of their similarities. R. siberu male then known Microhylidae is a group of small-sized frog which has no nuptial pad. The differences of their morphological has 14% number of total species in Sundaland. This character were predicted as the result of the difference family consists of ±6 genera (Calluella, between Sumatra and Peninsular Malaysia land mass Gastrophrynoides, Kalophrynus, Kaloula, Microhyla, and (Lim and Leong, 2004). Mentawai Island, Sumatra, and Metaphrynella) with ±43 of total species. Kalophrynus Peninsular Malaysia connected through Batu Island and Microhyla have the highest total species in Sundaland (Northern Siberu) (Heaney, 1986; Hall, 1996; Voris, compared with other genus. Most of Kalophrynus species 2000). The land mass difference allowed genetic flow are spread in Borneo, and some are spread in 3 other between Siberut, Sumatra, and Peninsular Malaysia. major islands (Das and Haas, 2003; Teynie et al., 2010; Rana siberu was geographically isolated and Chan et al., 2010; Das et al., 2007; Matsui, 2009). Each of experienced allopatric speciation. Simulation of Microhylidae genera experienced some speciation process geographical isolation was begun by the increase of which caused by many factors. seawater level that caused the ocean depth between Kalophrynus experienced some speciation processes Eastern Siberut and Western Sumatra reached 400 m which lead to the formation of new species, for example (Heaney, 1986; Hall, 1996; Voris, 2000). During the last Kalophrynus eok which found at high land on Kalabid, 250.000 years, sea surface was predicted to decreased into Serawak, and Borneo. It can be distinguished with other 120 m and caused the separation of Siberut and Batu Kallophrynus which found at other parts of Borneo by Island that resulted in high level of speciation and their morphological characters. K. eok is different from K. endemism (such as R. siberu) (Lim and Leong, 2004). baluensis which found at Sabah, Western Malaysia by the The other genus that experienced speciation is presence of weak subarticular tubercles on its toe and one Hylarana. H. rufipes from West Sumatra known as the subarticular tubercle located on its fifth toe. It is different result of sympatric speciation of H. chalconota based on with K. eok which has subarticular tubercle on its third Tjong et al. (2012) research. Recent studies found that toe. Beside that, K. baluensis have unwebbed toes which there are differences on the number of chromosomes different with K. eok that have toes webbed basally to between them. Djamhuriyah (2000), clarifies that H. below. Despite on its skin, K. baluensis has bright chalconota has 13 chromosome pairs which consists of 5 brownish color on dorsal, which differ with K. eok that pairs of large chromosome and 8 pairs of small has granul and maroon color on its dorsal (Kiew, 1994; chromosome. Whereas, H. rufipes has 6 pairs of large Das and Haas, 2003). Those differences are predicted as chromosome and 7 pairs of small chromosome, 1 pair the result of adaptation on their habitat where they live submetasentric and 12 pairs metasentric chromosome and lead to sympatric speciation between Kalophrynus in (Tjong et al., 2012). Borneo.

Bufonidae Megophrydae Bufonidae family (True Toads) consists of ±9 genera Megophrydae in Sundaland consists of ±5 genera (Ansonia, Bufo, Ingerophrynus, Duttaphrynus, (Leptobrachella, Leptobrachium, Leptolalax, Megophrys, Leptophryne, Nectophryne, Pedostibes, Pelophryne, and and Xenophrys) with ±37 species. Leptobrachium and Phrynoides) with ±53 species in Sundaland (Grismer, Megophrys are genera which contributed more species 2007; Grismer et al., 2006; Grismer, 2006; Marx, 1976; than the others (Teynie et al., 2010; Iskandar and Erdelen, Marx, 1958; Chan et al., 2010; Teynie et al., 2010; Wood 2006; Inger and Iskandar, 2005; Chan et al., 2010; et al., 2008; Dring, 1983; Inger et al., 2001). Bufonidae in Grismer et al., 2004; Inger et al., 1995; Matsui, 2006; Sundaland was dominated by genus Ansonia which Inger, 1966). Most of Leptobrachium spread on Borneo distributed in Malay Peninsula, Borneo, and Sumatra and least spreading on other main islands (Malaya (Marx, 1976; Matsui, 2006; Teynie et al., 2010). Limited Peninsular, Sumatra, and Java).

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The divergence time of Leptobrachium predicted decrease this genus is being isolated of geographic that occured on early Eocene, whilethe speciation occurred on caused allopatric speciation. middle Eocene, then it simultaneously occurred on last Eocene or early Oligocene (Matsui et al., 2010; Zhao and Dicroglossidae Morgan, 1985; Chung et al., 1998). Based on the Dicroglossidae in Sundaland consists of ±4 genea distribution, speciation of Leptobrachium predicted (Fejervarya, Limnonectes, Occidozyga, and Ingerana) happened in on late-Miocene or early-Pliocene, when with ± 33species. This family was dominated by Borneo has lost connection with its mainland (Lloyd, Limnonectes which distributed in four major islands of 1978 cited in Wilson and Moss, 1999; Matsui et al., Sundaland. Limnonectes became a good example for 2010). At that period of time, Peninsular Malaysia, speciation despite on its high morphological similarity Sumatra, and Java were connected with land, meanwhile and genetic differentiation on its species (Djong et al., Borneo was least isolated with sea (Inger and Voris; 2010). This genus consists of 2 species complex which 2001; Chappell & Shackleton, 1986). That event affected have high morphological character similarity between on genetic flow of the islands, they lost connection with each species, L. blythii and L. kuhlii. L. blythii was other main islands then allowed isolation for certain distributed in Myanmar, Thailand, Peninsular Malaysia, period of time. One of the example is the research of and Northern Sumatra only. This condition was predicted Matsui et al. (2010) which indicated that speciation of L. as the result of its lifestyle. Its reproduction occurs in low nigrops from its ancestor L. hendricksoni dan L. hasseltii stream river and related to ancient river system in with divergence time estimation at last-Oligocene until Northern Sumatra which connected to Peninsular last-Miocene (30-14 Mya before present). The speciation Malaysia and end in Southern China sea and Andaman has expected as the impact of land connection disruption sea. During Pleistocene period, Northern Sumatra has 2 between Borneo and Peninsular Malaysia which caused river systems, Malacca strait which flowed between isolation between Borneo and Malayan peninsular Malay Peninsula and Sumatra then ended in Andaman population. sea,Siam river which ended in Southern China Sea. The Megophrydae has wider distribution compared to river system of Malacca straits comprised river at Leptobrachium. It distributed on four main islands Northern Sumatra, Riau, and some at Peninsular (Peninsular Malaysia, Borneo, Sumatra, and Java). The Malaysia. While Siam river system comprised of Riau evenly distribution expected similar to Pliocene period and some at Peninsular Malaysia (Voris, 1153; Tjong et and Pleistocene until the last 10.000 year which on that al., 2010). Both rivers system did not connected to the period there were repeating increasing and decreasing of river system in Java. This condition allowed L. blythii to sea level (Woodruff & Turner, 2009). As the spread from Peninsular Malaysia to northern Sumatera consequence, when the increasing of sea level genus and to develop sympatric speciation was occurred Megophrys could make a move, while when sea level between L. blythii species.

es Number ofSpeci New

Figure 3. Total of new species discovered based on 6 main family (Bufonidae, Ranidae, Megophrydae, Microhylidae, Rhacophoridae, and Dicroglossidae) in 4 major islands (Malay Peninsular, Borneo, Sumatra, and Java) and adjacent Island since 1981 in a range of 10 years.

New Species in Sundaland level of amphibian which formed some new species in Zoogeographical history of Sundaland has drawn Sundaland. The total new species which found in interest of some researcher to study about it. It was Sundaland was approximately 51 species from family proven by some researchers about the high speciation Bufonidae, Ranidae, Megophrydae, Microhylidae,

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Rhacophoridae, and Dicroglossidae. The new species Microhylidae (14%), Megophrydae (12%), and which found in Sundaland was dominated by Dicroglossidae (11%). List of new species can be seen at Rhacophoridae (25%), Ranidae (20%), Bufonidae (18%), Table 1.

Table 1. New species which found in Sundaland at year range 1981 – 2015. No. Taxa Origin Bufonidae 1 Ansonia anotis (Inger et all., 2001) Borneo 2 Ansonia echinata (Inger and Stuebing, 2009) Borneo 3 Ansonia endauensis (Grismer, 2006) Peninsular Malaysia 4 Ansonia jeetsukumarani (Wood, Grismer, Ahmad, and Senawi, 2008) Peninsular Malaysia 5 Ansonia kraensis (Matsui et al., 2005) Peninsular Malaysia 6 Ansonia latiffi (Wood, Grismer, Ahmad & Senawi, 2008) Peninsular Malaysia 7 Ansonia torrentis (Dring, 1983) Borneo 8 Ingerophrynus gollum (Grismer, 2007) Peninsular Malaysia 9 Pelophryne api (Dring, 1983) Borneo 10 Pelophryne linanitensis (Das, 2007) Borneo 11 Pelophryne murudensis (Das, 2008) Borneo 12 Pelophryne rhopophilius (Inger and Steubing, 1996) Borneo 13 Pelophryne saravacensis (Inger and Stuebing, 2009) Borneo Ranidae 14 Amolops amoropalamus (Matsui, 1986) Borneo 15 Amolops macrophthalmus (Matsui, 1986) Borneo 16 Amolops orphnocnemis (Matsui, 1986) Borneo 17 Rana asperata (Inger et all., 1996) Borneo 18 Rana banjarana (Leong and Lim, 2003) Peninsular Malaysia 19 Rana rhacoda (Inger et all., 1996) Borneo Megophrydae 20 Leptobrachella palmate (Inger and Steubing, 1991) Borneo 21 Leptolalax kajangensis (Grimer et al., 2004) Peninsular Malaysia 22 Leptolalax maurus (Inger et all., 1997) Borneo 23 Megophrys edwardinae (Inger, 1989) Borneo 24 Megophrys dringi (Inger, 1995) Borneo Microhylidae 25 Calluella minuta (Das et al., 2004) Peninsular Malaysia 26 Kalophrynus calciphilus (Dehling, 2011) Borneo 27 Kalophrynus eok (Das and Haas, 2003) Borneo 28 Kalophrynus nubicola (Dring, 1983) Borneo 29 Kalophrynus yongi (Matsui, 2009) Peninsular Malaysia 30 Microhyla orientalis (Matsui, Hamidy, Eto, 2013) Java (adjacent island) 31 Microhyla maculifera (Inger, 1989) Borneo 32 Microhyla mantheyi (Das et al., 2007) Peninsular Malaysia Rhacophoridae 33 Philautus aurantium (Inger, 1989) Borneo 34 Philautus bunitus (Inger et all., 1995) Borneo 35 Philautus davidlabangi (Matsui, 2009) Borneo 36 Philautus disgregus (Inger) Borneo 37 Philautus erythrophthalmus (Steubing and Wong, 2000) Borneo 38 Philautus juliandringi (Dehling, 2010) Borneo 39 Philautus refugii (Inger and Steubing, 1996) Borneo 40 Polypedates chlorophthalmus (Das, 2005) Borneo 41 Rachophorus norhayatii (Chan and Grismer, 2010) Peninsular Malaysia 42 Rhacophorus achantharrhena (Harvey et al., 2002) Sumatra 43 Rhacophorus barisani (Harvey et al., 2002) Sumatra 44 Rhacophorus catamitus (Harvey et al., 2002) Sumatra 45 Rhacophorus gadingensis (Das and Haas, 2005) Borneo 46 Rhacophorus kajau (Dring, 1983) Borneo 47 Theloderma licin (Macleod and Ahmad, 2007) Peninsular Malaysia Dicroglossidae 48 Ingerana rajae (Iskandar, Bickford, Arifin, 2011) Borneo 49 Limnonectes cintalubang (Matsui, Nishikawa, Eto, 2014) Borneo 50 Limnonectes hikidai (Matsui and Nishikawa, 2014) Borneo

The total new species which found in Sundaland was explored Sundaland are Robert F. Inger, Masafumi estimated during 1981-2015 shown in Figure 3. The data Matsui, and Indraneil Das. The lack of new species shows that most of new species was found in Borneo. It discovery in Java and Sumatra probably was caused by probably caused by geological history of Borneo which the less concern from researchers. already explained in previous chapter. The highest discovery of amphibian new species was achieved at ZOOGEOGRAPHY OF SUNDALAND range of year 2001 – 2010 in Peninsular Malaysia, Borneo, and Sumatra with ±25 of total species. That Distribution of Anura (Family Bufonidae, Ranidae, probably caused by researcher interest on that year to Megophrydae, Microhylidae, Rhacophoridae, and explore Sundaland. Some of researchers which already Dicroglossidae) in four major islands of Sundaland were

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measured by the total number of spesies which spread in Java island has 59 young volcanoes with more than that four major islands. The result shows that Borneo has 2000 m in height and volcano from Tertiary period. One the highest number of total species, meanwhile Java is the of the most powerful supervolcanic eruption in Java lowest one (Figure 2). Species distribution of Anura in island was Krakatau in 1883 (Self and Rampino, 1981). four major islands, probably affected by amphibian That eruption covered Java island with lava and distribution at Plio-Pleistocene period (Wilting et al., dustwhich caused extinction of local species. In addition, 2012). Java island became drier and had poorer habitat for its Borneo has the highest number of total species (from organism compared to Sumatra and Borneo (Mayr, 1944). family Bufonidae, Ranidae, Megophrydae, Microhylidae, Rhacophoridae, and Dicroglossidae) compared to the three other islands. It was predicted to be related to CONCLUSION geological history of Borneo. Borneo was not included in Sundaland which located in equator line has the series of volcanic ring which related to the tectonic interesting geological history and topography. Therefore, plate. Series of volcanic ring was stretched from Sumatra its zoogeography, ecological barrier, and diversity have to Javawhich allowed supervolcanic eruptions which drawn interest of many researchers. Meanwhile, during resulted in the extinction of species (Mayr, 1944). In 1981-2015 most of researchers concern only on geological history, Borneo has no volcano-mountain row Peninsular Malaysia and Borneo. Therefore, further study which caused supervolcanic eruption and species about the exploration of amphibian in Sumatra and Java extinction. These cases caused the gene flow that goes to needed to increase amphibian database of Sundaland. Borneo is more stable and make the possibility the sympatric speciation to be occured. 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